a new key to the genera of liverworts of colombia · dumort., clevea lindb., isopaches buch,...

225

doi: https://dx.doi.org/10.15446/caldasia.v38n2.60915Caldasia 38(2): 225-249. 2016http://www.revistas.unal.edu.co/index.php/cal

INTRODUCTION

Colombia has a very rich liverwort flora. With about 715 accepted species, in 40 families, the country ranks second among the countries of tropical America, after Brazil with 725 species, and has more than half of the species recorded from tropical America (Gradstein & Uribe-M. 2016). Many species have been newly discovered in the country in recent years, and our knowledge of the liverwort flora of Colombia has been greatly improved by recent taxonomic revisions and monographs (e.g., Reiner-Drehwald & Goda 2000, Heinrichs 2002, Dauphin 2003, Váňa 2003, Bischler et al. 2005, Costa 2008, Uribe-M. 2008, Campos-Salazar & Uribe-M. 2012, Gradstein & Ilkiu-Borges 2015, Gradstein 2016, Ilkiu-Borges 2016). In spite of these studies and the availability of a modern catalogue (Gradstein & Uribe-M. 2016), identification of the liverworts of Colombia remains cumbersome. The key to the genera of liverworts of Colombia (Uribe-M. & Aguirre-C. 1997) is out of date

A new key to the genera of liverworts of Colombia

Nueva clave para los géneros de Hepáticas de Colombia

S. RobbeRt GRadSteinMuséum National d’Histoire Naturelle, Departement Systématique et Evolution, Paris, France, [email protected]

AbsTRACTA key is presented to 130 liverwort genera recorded from Colombia. Some genera found in neighboring countries, but not yet in Colombia, are also included in the key.

Key words. Colombia, hepatics, identification, taxonomy.

REsUMENSe presenta una clave taxonómica para los 130 géneros de hepáticas registrados en Colombia. Algunos géneros registradas de países vecinos, pero aún no de Colombia, también son incluidos.

Palabras clave. Colombia, hepáticas, identificación, taxonomía.

and keys to the species are very scattered in the literature and often difficult of access. The lack of a modern means of identification has prompted the author to prepare an identification manual for the liverworts of Colombia (Gradstein in prep.). As a first step, a treatment of the genus Plagiochila (Dumort.) Dumort., the largest genus of the liverworts, was published (Gradstein 2016) and a revision of the large and troublesome genus Bazzania Gray is in preparation. The present paper presents a new key to the genera of liverworts of Colombia. Although 132 genera were listed in the recent catalogue (Gradstein & Uribe-M. 2016), several taxonomic changes have occurred since the publication of the catalogue leading to modification of the total number of liverwort genera in Colombia. Aureolejeunea R.M. Schust. and Omphalanthus Lindenb. & Nees have been transferred to Cheilolejeunea (Spruce) Steph. (We et al. 2015), Austrofossombronia R.M.Schust. is now included in Fossombronia Raddi (Stotler

taxonomía y SiStemática

226

et al. 2016), and Prionocolea R.M.Schust. and Taxilejeunea (Spruce) Steph. are synonyms of Lejeunea Lib. (Gradstein et al. 2016). On the other hand, Archilejeunea subgen. Dibrachiella (Spruce) Schiffn. with three species in Colombia was raised to genus level (Shi et al. 2015), Cololejeunea minutissima (Sm.) Schiffn. was transferred to Myriocoleopsis Schiffn. (Yu et al. 2015), a rare rheophytic genus previously unknown in Colombia, and Platycaulis R.M.Schust. was newly recorded from Colombia (Gil-Novoa et al. 2015). As a result, 130 genera are currently accepted for Colombia and are included in the key. Some genera known from neighboring areas but not yet from Colombia (e.g., Anthelia (Dumort.) Dumort., Clevea Lindb., Isopaches Buch, Lobatiriccardia Furuki, Nanomarsupella R.M.Schust., Schusterolejeunea Grolle, Spruceanthus Verdoorn) are also included. Genera with only one species in Colombia as well as a few other taxa are directly keyed to species.

CLAssIFICATION

The classification of the genera into families is according to Gradstein & Uribe-M. (2016) except for Stephaniella and Stephaniellidium, which are placed in a separate family, Stephaniellaceae, following Schuster (2002); see also Juárez-Martínez et al. (2016). Classification of the families into orders, subclasses and classes follows Crandall-Stotler et al. (2009) except for Pleuroziaceae which are placed in a separate subclass, Pleuroziidae, following Frey (2009). Author citations of the genera follow Söderström et al. (2016) but the use of “ex” (e.g., Gottsche ex Steph.) is omitted and only the second, validating author (in this case Steph.) is cited since the use of “ex” in author citation is not obligatory (McNeill et al. 2012).

MARCHANTIOPSIDA

Aytoniaceae Asterella P.Beauv. Plagiochasma Lehm. & Lindenb.Corsiniaceae Cronisia Berk.Cyathodiaceae Cyathodium Kunze Dumortieraceae Dumortiera NeesLunulariaceae Lunularia Adans.Marchantiaceae Marchantia L.Monocleaceae Monoclea Hook.Ricciaceae Riccia L. Ricciocarpos CordaTargioniaceae Targionia L.

JUNGERMANNIOPSIDA

PELLIIDAEFossombroniaceae Fossombronia RaddiPallaviciniaceae Jensenia Lindb. Pallavicinia Gray Symphyogyna Nees & Mont.Pelliaceae Noteroclada Hook.f. & WilsonMETZGERIIDAEAneuraceae Aneura Dumort. Riccardia GrayMetzgeriaceae Metzgeria RaddiPLEUROZIIDAE Pleuroziaceae Pleurozia Dumort.JUNGERMANIIDAE1. JUNGERMANNIALES

Key to liverworts of Colombia

227

Gradstein

Acrobolbaceae Acrobolbus Nees Lethocolea Mitt. Tylimanthus Mitt.Adelanthaceae Adelanthus Mitt. Pseudomarsupidium HerzogArnelliaceae Gongylanthus Nees balantiopsaceae Isotachis Mitt. Neesioscyphus Grolle Ruizanthus R.M.Schust.Calypogeiaceae Calypogeia RaddiCephaloziaceae Alobiellopsis R.M.Schust. Cephalozia (Dumort.) Dumort. Fuscocephaloziopsis Fulford Nowellia Mitt. Odontoschisma (Dumort.) Dumort.Cephaloziellaceae Cephaloziella (Spruce) Schiffn. Cephaloziopsis (Spruce) Schiffn. Cylindrocolea R.M.Schust. Gymnocoleopsis (R.M.Schust.) R.M.Schust. Kymatocalyx Herzog Gymnomitriaceae Gymnomitrion Corda Marsupella Dumort.Herbertaceae Herbertus Gray Triandrophyllum Fulford & HatcherJamesoniellaceae Syzygiella SpruceJungermanniaceae Jungermannia L.Lepicoleaceae Lepicolea Dumort.Lepidoziaceae Bazzania Gray Kurzia G.Martens Lepidozia (Dumort.) Dumort. Micropterygium Gottsche Monodactylopsis (R.M.Schust.)

R.M.Schust. Mytilopsis Spruce Paracromastigum Fulford & J.Taylor Pseudocephalozia R.M.Schust. Pteropsiella Spruce Telaranea Schiffn. Zoopsidella R.M.Schust. Lophocoleaceae Clasmatocolea Spruce Cryptolophocolea L.Söderstr. Heteroscyphus Schiffn. Leptoscyphus Mitt. Lophocolea (Dumort.) Dumort. Platycaulis R.M.Schust.Plagiochilaceae Plagiochila (Dumort.) Dumort.Pseudolepicoleaceae Blepharostoma (Dumort.) Dumort. Chaetocolea Spruce Temnoma Mitt. scapaniaceae Anastrophyllum (Spruce) Steph. Diplophyllum (Dumort.) Dumort. Heterogemma (Jörg.) Konstant. & Vilnet Lophonardia R.M.Schust. Scapania (Dumort.) Dumort. Schistochilopsis (N.Kitag.) Konstant. Sphenolobus (Lindb.) Berggr.solenostomataceae Nardia Gray Solenostoma Mitt.stephaniellaceae Stephaniella J.B.Jack Stephaniellidium GrolleTrichocoleaceae Leiomitra Lindb. 2. PORELLALESFrullaniaceae Frullania RaddiJubulaceae Jubula Dumort. Lejeuneaceae Acanthocoleus R.M.Schust. Acrolejeunea (Spruce) Steph. Anoplolejeunea (Spruce) Schiffn. Archilejeunea (Spruce) Steph.

228

Blepharolejeunea S.W.Arnell Brachiolejeunea (Spruce) Schiffn. Bromeliophila R.M.Schust. Bryopteris (Nees) Lindenb. Caudalejeunea Schiffn. Ceratolejeunea (Spruce) J.B.Jack & Steph. Cheilolejeunea (Spruce) Steph. Cololejeunea (Spruce) Steph. Colura (Dumort.) Dumort. Cyclolejeunea A.Evans Dibrachiella (Spruce) X.Q.Shi et al. Dicranolejeunea (Spruce) Schiffn. Diplasiolejeunea (Spruce) Schiffn. Drepanolejeunea (Spruce) Steph. Frullanoides Raddi Fulfordianthus Gradst. Harpalejeunea (Spruce) Schiffn. Lejeunea Lib. Lepidolejeunea R.M.Schust. Leptolejeunea (Spruce) Steph. Lindigianthus Kruijt & Gradst. Lopholejeunea (Spruce) Steph. Luteolejeunea Piippo Marchesinia Gray Mastigolejeunea (Spruce) Steph. Metalejeunea Grolle Microlejeunea (Spruce) Steph. Myriocoleopsis Schiffn. Neurolejeunea (Spruce) Schiffn. Odontolejeunea (Spruce) Schiffn. Otigoniolejeunea (Spruce) Schiffn. Otolejeunea Grolle & Tixier Pictolejeunea Grolle Prionolejeunea (Spruce) Schiffn. Pycnolejeunea (Spruce) Schiffn. Rectolejeunea A.Evans Schiffneriolejeunea Verd. Stictolejeunea (Spruce) Schiffn. Symbiezidium Trevis. Thysananthus Lindenb. Verdoornianthus Gradst. Xylolejeunea X.L.He & GrollePorellaceae Porella L.Radulaceae Radula Dumort.

KEy TO THE GENERA

The terminology used in the key follows the Guide to the Bryophytes of Tropical America (Gradstein et al. 2001). Users of the key are advised to consult the latter publication, especially the glossary, descriptions and illustrations, which might help in clarifying difficulties that may arise.

Introductory key

1. Plants with stem and leaves ................. 21. Plants thalloid ...................................... 42. Leaves divided into segments ..................

................. Jungermanniopsida: Key 22. Leaves not divided into segments ........ 33. Leaves in 3 equal rows, leaf insertion

transverse. [Plants growing erect from a leafless rhizome, rhizoids absent]. Known from Ecuador, to be expected in Colombia... Haplomitrium blumei (Nees) R.M.Schust. (Haplomitriopsida)

3. Leaves not in 3 equal rows (leaf insertion various).... Jungermanniopsida: Key 2

4. Thallus with air chambers (cross section) and numerous small pores on dorsal surface. Thallus underside with scales, at least near the tip. Antheridia and archegonia usually born in receptacles ...................... Marchantiopsida: Key 1

4. Thallus lacking air chambers and pores. Thallus underside with or without scales ............................................................. 5

5. Upper surface of the thallus with a median groove. Plants often growing in rosettes (loosely forked in aquatic plants). Sporophytes embedded in the thallus ...................... Ricciaceae: Key 1

5. Upper surface of the thallus without median groove. Plants not growing in rosettes. Sporophytes not embedded in the thallus ........................................... 6

6. Rhizoids papillose. Thallus large, 1-3 cm wide, surface uniformily green. Gametangia produced in rounded


229

Gradstein

receptacles with blacks hairs at the margin...Dumortiera hirsuta (Sw.) Nees

6. Rhizoids smooth (exceptionally a few rhizoids with papillae present). Thallus small or large, surface variable, uniformly green or not. Gametangia not produced in receptacles with black hairs at the margin ....................................... 7

7. Thallus large, (0.5-)1-3 cm wide, without midrib, margins crispate-undulate, surfa-ce with numerous small whitish dots (fresh material). Antheridia produced in rounded or elongate receptacles on the thallus surface ....... Monoclea gottschei Lindb.

7. Thallus smaller, with or without midrib, margins plane or undulate, surface without small dots. Antheridia not produced in receptacles on the thallus surface ...... Jungermanniopsida: Key 2 (couplets 5-6)

Key 1. Marchantiopsida(complex thalloid liverworts)

1. Upper surface of the thallus with pores, appearing as numerous tiny, colorless or whitish dots. Air chambers present (cross-section) ..................................... 2

1. Thallus lacking pores. Air chambers absent or very inconspicuous ............ 12

2. Gemma receptacles present on dorsal surface of thallus ................................. 3

2. Gemma receptacles absent .................. 43. Gemma receptacles cup-shaped ............

............................................ Marchantia3. Gemma receptacles a lunate “scale” .....

........... Lunularia cruciata (L.) Dumort.4. Thallus with a median groove, often

growing in (partial) rosettes, sometimes floating on water. Sporophyte embedded in the thallus (Ricciaceae) ................... 5

4. Thallus lacking median groove, usually not in rosettes. Sporophyte not embedded in the thallus ....................... 6

5. Plants floating on water (rarely on wet soil). Pores present, bounded by a ring

of cells. Ventral scales large, ligulate, with oil cells. Dorsal thallus surface ± dark-green and reticulate ....................... ............ Ricciocarpos natans (L.) Corda

5. Plants on soil, rarely in water (Riccia stenophylla). Pores ± absent, when present without ring of cells. Ventral scales usually absent, when present small and without oil cells. Dorsal thallus surface pale green to glaucous green, usually not reticulate ........ Riccia

6. Thallus very thin, 2-layered, translucent. Ventral scales lacking (except at thallus apex). Sporophytes on the ventral thallus surface ............................... Cyathodium

6. Thallus thicker, not translucent. Ventral scales present throughout. Sporophytes not on the ventral thallus surface ........ 7

7. Air chambers without photosynthetic fi-laments. Female receptacles stalked..... 8

7. Air chambers with photosynthetic filaments. Female receptacles stalked or not stalked ......................................... 10

8. Pores surrounded by several rings of differentiated cells. Archegoniophore arising at the thallus apex on a 10-40 mm long stalk. Sporophyte surrounded by a large, conical, whitish or purplish pseudoperianth which hangs down beneath the receptacle and is split above into 8-16 narrow segments. Thallus glossy green to reddish ........... Asterella

8. Pores surrounded by only one ring of differentiated cells. Archegoniophore arising from the thallus surface (along the midline) on a very short, 1-5 mm long stalk. Sporophyte not surrounded by a large, conical, whitish or purplish pseudoperianth. Thallus light green to blue-green ........................................... 9

9. Thallus dull blue-green, surface not distinctly reticulate, pores incons-picuous. Ventral scales broadly ovate, entire, in 2 well-defined rows, each scale with 1-3 distinct, lanceolate appendages. Common species .............. Plagiochasma rupestris (Forster) Steph.

230

9. Thallus bright light green, surface conspicuously reticulate, pores distinct. Ventral scales lanceolate, ± toothed, in several ill-defined rows, each scale with an ill-defined, linear-lanceolate appendage. Known from Ecuador (but very rare), to be expected in Colombia .................................... Clevea spathysii (Lindenb.) Müll.Frib. (Cleveaceae)

10. Pores compound, barrel-shaped, formed of several layers of cells (cross-section). Male and female receptacles stalked and variously lobed .................. Marchantia

10. Pores simple, of one layer of cells. Receptacles sessile or only the female receptacle stalked .............................. 11

11. Thallus completely black when dry, margins and underside dark purplish-black. Ventral scales lanceolate, longer than wide. Gemmae lacking. Sporophyte in a dark, swollen, mussel-like involucre below the thallus apex ......................... ........................ Targionia hypophylla L.

11. Thallus not black when dry, margins and underside green or tinged with purple. Ventral scales very broad, much wider than long. Gemmae usually produced on thallus surface in a narrow, lunate receptacle. Sporophyte (very rare) on a stalked receptacle .................................. .......... Lunularia cruciata (L.) Dumort.

12. Plants small, thallus segments less than 0.5 cm wide, often in (partial) rosettes. Thallus surface often with a median groove ............................ (Ricciaceae) 5

12. Plants large, thallus segments 1-3 cm wide, never in rosettes. Thallus surface without median groove ..................... 13

13. Thallus surface with numerous small whitish dots. Thallus margins undulate-crispate. Thallus without midrib ............ ............................... Monoclea gottschei

13. Thallus surface without whitish dots. Thallus margins plane. Thallus with a narrow midrib formed by rhizoids on the ventral thallus surface ............................

............................... Dumortiera hirsuta

Key 2. Introductory key to Jungermanniopsida

1. Plants with stem and leaves ................ 21. Plants thalloid .................................... 52. Plants robust, to 10 cm long and 4-7

mm wide, reddish or purplish. Leaves deeply concave, ovate-oblong, strongly clasping the stem, undivided, leaf margin with a conspicous white margin. Underleaves absent. Perianth terete (or plicate).................................... Pleurozia

paradoxa (J.B.Jack) Schiffn. (Pleuroziidae)2. Plants different .................................... 33. Leaves undivided, succubous. Leaf base

several layers of cells thick. Cells thin-walled, without trigones. Rhizoids violet or colorless. Archegonia and antheridia on dorsal surface of stem. On soil or rock ............................. Pelliidae: Key 3

3. Leaves divided into segments or undivided, incubous, succubous or transverse. Leaf base only one layer of cells thick. Cells thin-walled or thick-walled, often with trigones. Rhizoids not violet. Archegonia on apex of stem or short branch, antheridia in leaf axils. On bark, living leaves, rotten wood, rock and soil (Jungermanniidae – foliose liver worts) ........................................... 4

4. Leaves with a ventral lobule (= small or large fold or sac appressed to the leaf, on the ventral side of leaf), incubous ......... ................................. Porellales: Key 11

4. Leaves without lobule ........................... ...................... Jungermanniales: Key 4

5. Thallus unbranched or with a few short, ventral-intercalary branches. Thallus margins, at least near the apex, with long, sausage-shaped slime papillae. Archegonia and antheridia on leafy branches .................................. Pterop -siella serrulata Steph. (Lepidoziaceae)

5. Thallus variously branched. Thallus margins without sausage-shaped slime papillae. Archegonia and antheridia on


231

Gradstein

the thallus surface or in cavities, not on leafy branches ..................................... 6

6. Thallus midrib with a central strand. Gametoecia on the dorsal side of the midrib ........... Pallaviciniaceae: Key 3

6. Thallus midrib without central strand, or midrib absent. Gametoecia on the ventral side of the midrib or at the thallus margin ................. Metzgeriidae: Key 3

Key 3. Metzgeriidae and Pelliidae (simple thalloid liverworts)

1. Plants with stem and leaves (Pelliidae). 21. Plants thalloid (Metzgeriidae and Pellii-

dae) ..................................................... 32. Rhizoids pale brown. Plants 4-10 mm

wide, glaucous-green or deep green. Leaf margins flat, entire ................................

............... Noteroclada confluens Taylor2. Rhizoids purplish. Plants 2-4(-5) mm

wide, pale-green, usually forming rosettes. Leaf margins undulate, entire or toothed....................... Fossombronia

3. Thallus simple or pinnate, without midrib or with midrib only on branches. Gametoecia on the thallus margin or on short lateral branche...... (Aneuraceae) 4

3. Thallus simple or dichotomous, with a midrib throughout. Gametoecia on the midrib (dorsal or ventral side).............. 6

4. Thallus simple or scarcely branched, more than 2 mm wide, prostrate. Oil bodies 10-40 per cell. Male branches with antheridia in 2-6 rows........ Aneura

4. Thallus distinctly branched (irregularly to regularly pinnate), 0.5-2(-8) mm wide, prostrate or erect. Oil bodies less than 10 per cell. Male branches with antheridia always in 2 rows................. 5

5. Thallus 0.5-2 mm wide, thallus 1-3-pinnate with short or long branches. Archegonia on short lateral branches, without paraphyses................. Riccardia

5. Thallus 2-8 mm wide, thallus 1-pinnate with rather short and wide branches (branches ca. 1-2× as long as wide).

Archegonia in incisions of the thallus margin surrounded by numerous crowded, multicellular paraphyses, not on branches. Known from Ecuador, to be expected in Colombia.... Lobatiriccardia

6. Thallus 0.5-2(-3) mm wide, margins with unicellular hairs (= rhizoids). Midrib without central strand (cross section). Gametoecia on the ventral side of the midrib.......................... Metzgeria

6. Thallus usually more than 3 mm wide, margins without hairs. Midrib with central strand. Gametoecia on the dorsal side of the midrib (Pallaviciniaceae) 7

7. Thallus divided into lobes...................... ........................................ Symphyogyna

7. Thallus not lobed................................. 88. Thallus ascendent to erect from a

prostrate rhizome................................. 98. Thallus prostrate or slightly ascendent,

without prostrate rhizome.................. 109. Archegonia surrounded by a ring of

scales forming a cup-like involucre. After fertilization, a tubular pseudoperianth several times longer than the involucre developing from within the involucre.... ................................................. Jensenia

9. Archegonia merely with a small scale inserted behind them (= side directed to the base of the thallus), cup-like involucre and pseudoperianth lacking.... ........................................ Symphyogyna

10. Thallus margins with scattered slime hairs 2-4 cells long. Midrib with 1 central strand. Archegonia surrounded by a ring of scales forming a cup-like involucre. After fertilization, a tubular pseudoperianth several times longer than the involucre developing from within the involucre................................

........... Pallavicinia lyellii (Hook.) Gray10. Thallus margins without slime hairs.

Midrib with 1-3 central strands. Archegonia merely with a small scale inserted behind them (= side directed to the base of the thallus), cup-like

232

involucre and pseudoperianth lacking.... ........................................ Symphyogyna

Key 4. Introductory key toJungermanniales (foliose liverworts I)

1. Leaves deeply divided into hair-like segments which are not more than 1-2 cells wide at base................................. 2

1. Leaves undivided or divided into segments which are more than 2 cells wide at base.......................................... 4

2. Leaves divided to the base into 1-4 filaments (= uniseriate segments), margins entire. Plants very small, alga-like......................................... Telaranea

2. Leaves divided into 4-9 segments, margins strongly ciliate. Plants larger, not alga-like......................................... 3

3. Leaves 4-9-lobed, the segments split into numerous ciliate parts, the plants strongly “hairy”. Leaf cells with thin or uniformly thickened walls, trigones lacking. Underleaves smaller than lateral leaves. Plants whitish-green or brown ... .............. Leiomitra (= Trichocolea p.p.)

Probably all Trichocolea species from Colombia belong to Leiomitra but some poorly known species that are only known from sterile material have not yet been transferred to Leiomitra.

3. Leaves 4-lobed, the segments not split into numerous ciliate parts. Leaf cells with large trigones. Underleaves as large as lateral leaves. Plants gray-green or brown ......................................

Blepharostoma trichophyllum (L. )Dumort.4. Small whitish, worm-like plants with

densely imbricate leaves on soil in páramo................................................. 5

4. Plants different..................................... 75. Leaves undivided, opposite or alternate.

Leaf cells conspicuously elongate near the ventral margin........................................

............................ Arnelliaceae: Key 4b5. Leaves bifid, alternate. Leaf cells not

conspicuously elongate near the ventral margin.................................................. 6

6. Plants with 3 rows of leaves, moss-like. Perianth large, deeply plicate. Minute whitish, prostrate liverwort from superpáramo of Ecuador, to be expected in Colombia............................. Anthelia

juratzkana (Limpr.) Trevis. (Antheliaceae)6. Plants with 2 rows of leaves (underleaves

lacking). Perianth lacking or very small, inconspicuous.........................................

.................... Gymnomitriaceae: Key 77. Leaves with (1-)2 large, sausage-shaped

slime papillae at the tips of the segments or on the rounded margins. Plants very small, less than 1 mm wide, whitish-green, the leaves almost longitudinally inserted on the stem. On rotten wood, soil and rock in rainforest... Zoopsidella

7. Leaves without large slime papillae at the tips or on the margins..................... 8

8. Leaves incubous................................... 98. Leaves succubous or transverse......... 169. Underleaves and leaves of similar size,

bifid or trifid.............. (Herbertaceae) 109. Underleaves smaller than leaves........ 1110. Leaves 2-3-lobed, without vitta. Leaf

cells thin-walled to evenly thick-walled, ± without trigones................................... ............... Triandrophyllum subtrifidum (Hook.f. & Taylor) Fulford & Hatcher

10. Leaves bifid, with a broad vitta of elongate cells. Leaf cells with large trigo-nes........................................ Herbertus

11. Leaves with 3 or more lobes or teeth..... ............................................... Lepidozia

11. Leaves with 0-2 lobes or teeth........... 1212. Rudimentary lobule (consisting of a

few cells) present at ventral base of leaf. Stems fragile, ventral stem surface only 2-4 cells wide, hyalodermis often present. Ventral branches absent .................................................................Lejeu-neaceae (with reduced lobules): Key 12

12. Lobules completely lacking. Stems rigid, ventral stem surface more than 4


233

Gradstein

cells wide, hyalodermis absent. Ventral branches frequently present............... 13

13. Plants densely pinnate, growing upright on twigs near running water, with numerous clustered perianths on short lateral branches. Underleaves minute. Known from Ecuador, in rivers with granite rocks and strongly fluctuating water levels, between 1200-1800 m, to be expected in Colombia.........................

...................................... Colura irrorata (Spruce) Heinrichs et al. (Lejeuneaceae)13. Plants forked, irregularly branched or

unbranched, not growing on twigs near running water, perianths lacking or on ventral side of stem (not clustered on short lateral branches). Underleaves well-developed................................... 14

14. Stems forked, flagelliform ventral branches present. Leaf apex bifid or trifid (rarely entire).................. Bazzania

14. Stems simple or irregularly branched, flagelliform ventral branches lacking. Leaf apex entire or bifid.......................... .............................. (Calypogeiaceae) 15

15. Underleaves entire or short bifid. Underleaf cells very different from leaf cells (except in M. nephrostipa), conspicuously elongate, 2-5× as long as wide, without chlorophyll, hyaline. Plants deep green to brown. Leaf apex entire. Oil bodies brownish, finely granular. Vegetative reproduction by caducous leaves.................... Mnioloma

15. Underleaves bifid. Underleaf cells similar to leaf cells, 1-2× as long as wide, with chlorophyll, not hyaline. Plants pale green to bluish green. Leaf apex usually bifid. Oil bodies colorless or sepia, coarsely granular. Vegetative reproduction by gemmae produced on upright, flagelliform branches.................

............................................. Calypogeia16. Underleaves present................... Key 4a16. Underleaves absent (or very small)........ ................................................... Key 4b

Key 4a. Jungermanniales - Leaves succubous or transverse,

underleaves present

1. Leaf margins toothed........................... 21. Leaf margins entire.............................. 62. Leaves 4-lobed, underleaves as large as

leaves..................................... Lepicolea2. Leaves not 4-lobed, underleaves smaller

than leaves........................................... 33. Leaves folded and usually keeled, at

least above, the keel often winged.......... ........................... Lepidoziaceae: Key 83. Leaves not folded, without winged keel ............................................................. 44. Dorsal leaf base decurrent. Leaves

asymmetrical, ventral margin arched, dorsal margin ± straight. Stems brown (or bluish), with thick-walled cortex...... ........................................... Plagiochila

4. Dorsal leaf base not decurrent. Leaves ± symmetrical. Thick-walled cortex lacking................................................. 5

5. Leaves ± transversely inserted, deeply concave. Branches originating from the ventral side of the stem........... Isotachis

5. Leaves ± longitudinally inserted, not deeply concave. Branches originating from the lateral and ventral side of the stem................. Lophocoleaceae: Key 9

6. Leaves undivided................................. 76. Leaves divided into lobes.................. 107. Leafy plants at the base arising from a

leafless, stoloniform shoot...................... ........................... Lepidoziaceae: Key 87. Leafy plants not arising from a leafless,

stoloniform shoot................................. 88. Ventral stolons present Leaf insertion

reaching the dorsal midline of stem or not. Rhizoids scattered.. Odontoschisma

8. Ventral stolons lacking. Leaf insertion reaching the dorsal midline of stem. Rhizoids scattered or in bundles.......... 9

9. Underleaves entire, oblong-lanceolate. Rhizoids scattered. Plants green to reddish-brown or purple......................... ....... Nardia succulenta (Lehm.) Spruce

234

9. Underleaves bifid or toothed, rarely entire. Rhizoids in bundles. Plants green to brown.......... Lophocoleaceae: Key 9

10. Leaves divided into 3-5 lobes. Plants small, less than 2 mm wide, leafless stolons present or lacking................... 11

10. Leaves divided into 2(-3) lobes. Plants small or large...................................... 14

11. Underleaves very different from leaves, much smaller and only 2-lobed. [Leaves strongly concave, 4-5-lobed, sharply piliferous. On soil in superpáramo of the Sierra Nevada de Santa Marta].........

..Ruizanthus venezuelanus R.M.Schust.11. Underleaves similar to leaves (but

sometimes smaller)............................ 1212. Leaves succubous. [Leaves palmate,

divided to ca. 1/2 into narrow lanceolate, conspicuously diverging lobes]..............

................. Chaetocolea palmata Spruce12. Leaves transverse............................... 1313. Plants pale green, shallowly divided (to

1/4 or less) into 3-4 triangular lobes. Stem base stoloniform. Leaf cells thin-walled, cuticle smooth or sligthly papillose. Perianth on a short ventral branch, 3-keeled. In wet páramo....................

............................................... Pseudoce- phalozia quadriloba (Steph.) R.M.Schust.13. Plants brownish, deeply divided (to

3/4) into 3-4 stiff, subulate lobes. Stem base not stoloniform. Leaf cells thick-walled, cuticle strongly striate-papillose. Perianth on a long shoot. On rotten wood in upper montane cloud forest............ .....Temnoma chaetophylla R.M.Schust.

14. Leaves in 3 ± equal rows, underleaves about as large as leaves...................... 15

14. Leaves not in 3 equal rows, underleaves smaller than leaves............................. 17

15. Leaves distinctly elongate, (1.5-)2-7 times as long as wide. Leaves with a broad vitta of strongly elongate cells. Trigones very large................ Herbertus

15. Leaves shorter. Vitta absent. Trigones small or lacking.................................. 16

16. Plants very small, less than 1 mm wide, creeping, whitish in color. Leaves scarcely wider than the stem. On soil in superpáramo (above 4000 m). Known from Ecuador, to be expected in Colombia............... Anthelia juratzkana

16. Plants larger, creeping or upright, reddish, purple-brown or black. Leaves much wider than the stem. In water in páramo.............. Isotachis obtusa Steph.

(= Isotachis lacustris Herzog syn. nov.)17. Leaves transverse, strongly concave.

Sporophyte in a fleshy perigynium......... ................................................. Isotachis17. Leaves succubous. Sporophyte in a pe ri-

anth.................................................... 1818. Leaves and underleaves divided to the

middle into 2-3 narrow lobes (some 3-lobed leaves always present). Rhizoids in bundles................. Paracromastigum

18. Leaves and underleaves 2-lobed or undivided, never 3-lobed, less deeply divided. Rhizoids scattered or in bundles............................................... 19

19. Leaf insertion line not reaching dorsal midline of stem, dorsal side of stem “leaf-free”. Rhizoids scattered................

......................... Cephaloziaceae: Key 519. Leaf insertion line reaching dorsal

midline. Rhizoids in bundles.............. 2020. Plants often with some reddish

coloration (especially in the rhioids). Branching purely ventral. Leaf cells narrowly rectangular, cuticle with elongate papillae (“striate-papillose”)....

....................................... Neesioscyphus20. Plants without any reddish coloration.

Branching lateral and ventral. Leaf cells shorter, cuticle smooth or with small rounded papillae......................................

........................ Lophocoleaceae: Key 9


235

Gradstein

Key 4b. Jungermanniales - Leaves succubous or transverse, underleaves

absent or minute

1. Plants densely pinnate, growing upright on twigs near running water, with numerous clustered perianths on short lateral branches, perianth beaked. Underleaves minute. Known from Ecuador in rivers with granite rocks and strongly fluctuating water levels, between 1200-1800 m, to be expected in Colombia................................................

............. Colura irrorata (Lejeuneaceae)1. Plants different..................................... 22. Plants worm-like, often whitish, with

densely imbricate and appressed leaves, less than 2 mm wide. Leaves transverse, entire or bifid. On soil or rock in páramo................................................. 3

2. Plants not worm-like, never whitish, small or large, leaves less densely imbricate.............................................. 6

3. Leaves bifid, alternate. Leaf cells not conspicuously elongate near the ventral margin......... Gymnomitriaceae: Key 7

3. Leaves undivided, opposite or alternate. Leaf cells conspicuously elongate near the ventral margin................................ 4

4. Leaves opposite, the dosal leaf bases united, not falcate. Stems without paraphyllia. Stolons lacking. Plant green to brown to purplish or whitish...............

........................................ Gongylanthus4. Leaves alternate, the dorsal leaf bases

not united, usually falcate. Stems on the dorsal side with green paraphyllia. Stems attached to soil by long stolons. Plants grayish to pale brown................ 5

5. Leaves plicate, with some longitudinal folds. Paraphyllia lanceolate. Sporophyte produced in a marsupium........... Stepha-

niellidium sleumeri (Müll.Frib.) Grolle5. Leaves smooth, without folds. Para-

phyllia linear. Marsupium lacking, sporophyte surrounded by connate bracts

........................................... Stephaniella

6. Plants minute, less than 0.6 mm wide. Leaf cells usually small, 8-25(-30) µm, without trigones.......................................

..................... Cephaloziellaceae: Key 66. Plants larger. Cells with or without trigo-

nes........................................................ 77. Leaves opposite, dorsal leaf bases

united................................................... 87. Leaves alternate, dorsal leaf bases not

united................................................. 108. Plant creeping on soil, densely attached

to the soil with rhizoids. Leaf cells conspicously elongate towards the ventral leaf margin, trigones very small or lacking........................ Gongylanthus

8. Plant not creeping on soil, not densely attached to the soil. Leaf cells not conspicously elongate towards the ventral leaf margin, trigones usually lar ge..................................................... 9

9. Plants with reddish or purple pigmen-tation. Perianth inflated over its whole length, with several plicae..... Syzygiella

9. Plants lacking reddish or purple pigmentation. Perianth flattened towards the mouth, with 2 keels......... Plagiochila

10. Dorsal margin of leaves reflexed, dorsal leaf base distinctly decurrent. Leaf surface convex near dorsal margin and concave towards ventral margin......... 11

10. Dorsal margin of leaves plane or inflexed, not or little decurrent. Leaf surface ± plane or concave................. 13

11. Plants with reddish or purple pigmen-tation. Perianth inflated over its whole length .................................... Syzygiella

11. Plants lacking reddish or purple pigmentation. Perianth flattened towards the mouth or absent and replaced by a marsupium......................................... 12

12. Plants whitish-green or glaucous when fresh (gray or brown when dry). Leaf apex obliquely truncate-emarginate. Stems pale. Oil bodies brown, numerous, filling cell-lumen. Sporophyte in a marsupium.............................. Tyliman- thus laxus (Lehm. & Lindenb.) Spruce

236

12. Plants green or brown. Leaf apex various but usually not obliquely truncate-emarginate. Stems brown, darker than leaves. Oil bodies colorless, not filling cell-lumen. Sporophyte in a flattened perianth............................... Plagiochila

13. Leaves several layers of cells thick near the base........................ Pelliidae: Key 3

13. Leaves only one layer of cells thick.. 1414. Leafy plants arising from a creeping,

stolon-like shoot................................. 15 14. Leafy plants not arising from a creeping,

stolon-like shoot................................. 2015. Leaves transverse. [Dorsal margin

of leaves ± inflexed (towards stem). Stem rigid, with thick-walled cortex. Gametoecia at stem base on abbreviated branches] (Adelanthaceae)................. 16

15. Leaves succubous.............................. 1716. Cells in the upper part of the leaf ±

evenly thick-walled, without distinct trigones. Leaf base with a short vitta. Stem cortex well-developed, brown, of 2-3 cell layers (cross-section).................

............................................ Adelanthus16. All leaf cells with distinct trigones, cells

never evenly thick-walled. Vitta lacking. Stem cortex poorly developed, pale-colored, of 1 layer of cells......................

............................. Pseudomarsupidium 17. Leaf cells small, less than 25 µm,

without trigones. Plant less than 1 mm wide.................................. Kymatocalyx

17. Leaf cells larger, with or without trigones. Plants usually wider than 1 mm (rarely less than 1 mm)...................... 18

18. Leaf insertion line not reaching dorsal midline of stem, dorsal side of stem leaf-free. Plants frequently with upright flagelliform branches producing gem-mae.................. Cephaloziaceae: Key 5

18. Leaf insertion line reaching dorsal midline of stem, dorsal side of stem not leaf-free. Plants without such gemmi pa-rous branches..................................... 19

19. Leaf cells papillose. Plants whitish-green to yellowish-green (sometimes

brown when dry). Sporophyte in a marsupium............... Tylimanthus laxus

19. Leaf cells smooth. Plants green to reddish-brown to black. Sporophyte in a perianth.................................. Syzygiella

20. Leaves entire...................................... 2120. Leaves 2(-3)-lobed............................. 2421. Leaf cells conspicuously elongate

towards the ventral margin. [Plants whitish-green to yellowish-green. Lea-ves tongue-shaped. Sporophyte in a marsupium].............................................

Lethocolea glossophylla (Spruce) Grolle21. Leaf cells not elongate towards the

ventral margin. Sporophyte in a perianth.............................................. 22

22. Ventral stolons present. Leaf insertion reaching dorsal midline of stem of stem or not. Gemmae sometimes present. Gynoecia on a short ventral branch........

...................................... Odontoschisma22. Ventral stolons absent. Leaf insertion

always reaching dorsal midline of stem of stem. Gemmae absent. Gynoecia terminal on the main stem or on a long branch................................................ 23

23. Plants dark green to blackish-green, growing on wet rock in rivers at high elevations, rather large, 1.5-2 mmm wide, leaves longer than wide. Leaf cells small, less than 30 µm long in mid-leaf, thin-walled, trigones lacking or very small. Rhizoids few, not in bundles. Perigynium lacking (bracts not attached to perianth base)........................ Junger-

mannia ovatotrigona (Steph.) Grolle23. Plants light green to reddish or brown,

not growing on wet rock in rivers. Leaf cells (20-)30-60 µm long in mid-leaf, trigones present or lacking. Rhzoids often numerous, sometimes in bundles. Perigynium present (inner female bracts attached to perianth base) Solenostoma

24. Stem with a hyalodermis......................... ......................... Cephalo ziaceae: Key 524. Stem without hyalodermis................. 25


237

Gradstein

25. Leaves on sterile and male shoots with a very narrow base, the leaves 2-3× wider in the middle than at the base, caducous

............................................. Acrobolbus cuneifolius (Steph.) Briscoe (= Acrobol-

bus caducifolius R.M.Schust. syn. nov.)25. Leaves without very narrow base, not

caducous............................................ 2626. Cuticle smooth.. scapaniaceae: Key 1026. Cuticle finely papillose...................... 2727. Plants glossy whitish-green to light

brown (never reddish or purple). Gemmae absent. Oil bodies brown.........

............................................. Acrobolbus27. Plants greenish-brown to reddish or

purple. Gemmae present or absent. Oil bodies colorless.......................................

.......................... scapaniaceae: Key 10

Key 5. Cephaloziaceae

1. Leaves undivided. Stem without hyalodermis. Ventral stolons usually present.................................................. 2

1. Leaves shallowly or deeply bifid. Stem with hyalodermis. Ventral stolons lacking................................................. 3

2. Cells in midleaf 40-100 µm long, cell walls without trigones. Plants less than 1 mm wide. On bare clay soil below 2000 m.............................................. Alobiell-

opsis dominicensis (Spruce) Fulford2. Cells in midleaf 15-30(-40) µm long,

trigones usually present (rarely lacking). Plants (0.5-)1-2 mm wide. On rotten wood, humus or rock, from sea level to páramo.......................... Odontoschisma

3. Leaf base forming an inflated sac........... ................................................. Nowellia3. Leaf base not forming a sac................. 4 4. Leaves nearly longitudinally inserted,

dorsal leaf base decurrent. Leaf apex shallowly retuse or bifid to 1/6-1/3 .............................. Fuscocephaloziopsis

4. Leaves subtransversely inserted, dorsal leaf base not decurrent. Leaf apex bifid to 1/3-1/2............................. Cephalozia

Key 6. Cephaloziellaceae

1. Leaves undivided to retuse. Leafy stems erect, arising from creeping stolons........

.......................................... Kymatocalyx1. Leaves bifid to 1/3 or more. Leafy stems

creeping, stolons lacking...................... 22. Leaves obcuneate, widest above, apices

rounded. Plants light green, without any pigmentation, minute, ca. 0.5 mm wide..

.......................................Cephalo-ziopsis intertexta (Gottsche) R.M.Schust.

2. Leaves not obcuneate, apices acute or narrowly obtuse. Plants green to brown or reddish............................................. 3

3. Plants ca. 1 mm wide. Leaves distinctly concave, apices obtuse. Perianth terete. Only in páramo........... Gymnocoleop-sis cylindriformis (Mitt.) R.M.Schust.

3. Plants smaller, 0.3-0.6 mm wide. Leaves not concave, apices acute (rarely obtuse). Perianth plicate. Lowland, montane or páramo................................................. 4

4. Leaves subtransverse, hardly wider than the stem. Cells in midleaf very small, 8-15 µm, thick-walled (rarely thin-walled). Leaf insertion reaching the dorsal midline of the stem. Occurring between ca. 500-4000 m... Cephaloziella

4. Leaves distinctly succubous, distinctly wider than the stem. Cells in midleaf larger, 15-25 µm, thin-walled. Leaf insertion not reaching the dorsal midline of the stem. Below 1500 m.. Cylindricola

Key 7. Gymnomitriaceae

1. Plants minute, filiform, 0.1-0.2(-0.35) mm wide, creeping............................... 2

1. Plants larger, ascending to erect........... 32. Cells of leaves and stem epidermis

coarsely papillose (one large, rounded papilla per cell). Plants usually pale-colored. In rather dry superpáramo. Known from Venezuela and Ecuador, to be expected in Colombia.........................

Nanomarsupella xenophylla R.M.Schust.

238

2. Cells of leaves and stem epidermis smooth or somewhat mamillose, papillae lacking. Plants purplish to blackish........

............. Gymnomitrion atrofilum Váňa3. Leaves strongly concave from base

to apex (flattening of leaf impossible without tearing the leaf), very shallowly bifid to 1/8-1/5, with a broad, lunulate sinus and short, blunt tips.......... Gymno-

mitrion truncatoapiculatum Herzog3. Leaves concave only in the lower half,

flattened towards the apex, more deeply bifid, sinus rounded to acute (not broadly lunulate)............................................... 4

4. Plants whitish to pale brown. Perianth lacking............................ Gymnomitrion

4. Plants green to brown to reddish, not whitish. Perianth present but very small, hidden between the bracts... Marsupella

Key 8. Lepidoziaceae

1. Plants consisting of a small thallus with a narrow midrib and unistratose wings. Amazonia............ Pteropsiella serrulata

1. Plants not thalloid................................ 22. Plants consisting of minute shoots (less

than 0.2 mm wide), 2 cells wide, with rudimentary leaves, each leaf consisting of 1-2 cells with a slime papilla on top. In lowland rainforest.......... Monodacty-

lopsis monodactyla (Spruce) R.M.Schust.2. Plants larger, leaves not rudimentary,

made up of more than 2 cells............... 33. Leaves hair-like, consisting of 1-4

uniseriate filaments, undivided, leaf lamina lacking....................... Telaranea

3. Leaves not hair-like, with a short or long lamina.................................................. 4

4. Leaf cells very large, ca. 60-80 µm long in midleaf, thin-walled, without trigones. Leaf apex rounded to short-bifid, frequently with 1-2 large, sausage-shaped slime papillae (rarely without)...

............................................ Zoopsidella4. Leaf cells smaller, usually with trigones.

Leaf apex acute or divided into several

teeth or lobes, without sausage-shaped papillae................................................ 5

5. Leaves incubous.................................. 65. Leaves succubous or transverse.......... 76. Leaves divided into 4-6 lobes. Branching

(bi)pinnate. Ventral flagellae usually absent. Plants 0.5-1.5 mm wide..............

............................................... Lepidozia6. Leaves with 2-3 teeth at apex or

undivided. Branching forked or ventral. Ventral flagellae common. Plants 1-6 mm wide................................. Bazzania

7. Leaves folded and usually keeled, at least above, the keel often winged....... 8

7. Leaves not folded, 2-4-lobed............... 98. Leaf apex bifid to 1/4. Underleaves

absent......... Mytilopsis albifrons Spruce8. Leaf apex undivided or very short-bifid.

Underleaves present (sometimes very small)........................... Micropterygium

9. Leaves and underleaves shallowly divided (to 1/4 or less) into 3-4 triangular lobes. Plants growing upright from a stoloniferous base. In páramo.................

............... Pseudocephalozia quadriloba9. Leaves and underleaves more deeply

divided. Plants creeping or ascending, stoloniferous stem base present or absent................................................. 10

10. Stem leaves 2-3-lobed. Stems with a stoloniferous base..... Paracromastigum

10. Stem leaves 4-lobed. Stems without stoloniferous base....................... Kurzia

Key 9. Lophocoleaceae

1. Leaf surface covered by numerous small, spinose teeth................................. ...... Lophocolea muricata (Lehm.) Nees

1. Leaf surface smooth............................. 22. Leaves apex entire................................ 32. Leaf apex toothed ............................... 53. Leaves broadly reniform, almost twice

as wide as long, stiffly appressed to each other. Plants brown. In páramo bogs, rare... Platycaulis renifolia R.M.Schust.


239

Gradstein

3. Leaves ovate-orbicular to oblong, as wide as long or longer than wide, appressed or spreading......................... 4

4. Underleaves variable, entire to bifid on the same stem and usually narrower than the stem. Plants pale green, small, less than 1.5 mm wide. Leaf cells thin-walled, ± without trigones. On moist soil and rock in or near running water.. Clas-

matocolea vermicularis (Lehm.) Grolle4. Underleaves not entire and bifid on the

same stem. Plants green to brown, small or large. Leaf cells usually with trigo-nes.................................... Leptoscyphus

5. Leaves ± transverse, strongly caducous. Plants minute, yellowish green, ca. 0.5 mm wide. Tiny epiphyte in upper montane cloud forest and páramo.... Lo-

phocolea fragmentissima R.M.Schust.5. Leaves distinctly succubous, not

caducous. Plants different.................... 66. Underleaves free from the leaves......... 76. Underleaves connate with leaf bases on

one or both sides .................................. 87. Leaf apex bifid.................... Lophocolea 7. Leaf apex with 3 or more cilia (rarely

only 2)............................. Leptoscyphus 8. Plants fertile......................................... 98. Plants sterile....................................... 119. Gametangia on very short ventral

branches, hidden under the leaves. Perianth without keels.... Heteroscyphus

9. Gametangia on long branches, not hidden under the leaves. Perianth 2-3- keeled ................................................ 10

10. Leaves bifid............... Cryptolophocolea10. Leaves with 4-15 cilia, not bifid. Leptos- cyphus trapezoides (Mont.) L.Söderstr.11. Underleaves attached to leaves on one

side only. Leaf apex entire or with 2-3 obtuse lobes.................. Heteroscyphus contortuplicatus (Nees & Mont.) Grolle

11. Underleaves attached to leaves on both sides. Leaf apex with (0-)2-4 sharp teeth ........................................................... 12

12. Leaf apex with 2 teeth. Underleaves with 2-6 teeth or cilia (rarely more)........

.................................. Cryptolophocolea12. Leaf apex with 3 or more teeth or cilia.

Underleaves with numerous cilia....... 1313. Underleaves small, 2-3× wider than

the stem, subquadrate, deeply bifid. Dioicous....... Leptoscyphus trapezoides

13. Underleaves larger, 3-5× wider than the stem, reniform, not or shallowly bifid. Monoicous or dioicous... Heteroscyphus

Key 10. scapaniaceae

1. Leaves divided into a small dorsal lobe and a large ventral lobe........................ 2

1. Leaves not divided into a small dorsal lobe and a large ventral lobe. Mostly in páramo................................................. 3

2. Ventral leaf-lobe 2× longer than wide. Plants small, 1-1.5 mm wide, creeping on soil. Perianth inflated, plicate.............

........................................ Diplophyllum 2. Ventral leaf-lobe 1-1.5× longer than

wide. Plants 1.5-5 mm wide. Perianth flat, smooth............................. Scapania

3. Leaf margins on sterile stems toothed. Leaf cells large, 30-60 µm in midleaf, thin-walled, with very small trigones. . 4

3. Leaf margins on sterile stems entire. Leaf cells smaller................................. 6

4. Ventral stolons present..... Lophonardia4. Ventral stolons absent.......................... 55. Leaf cells very large, 40-60 µm in

midleaf. Stem epidermis cells 4-8× longer than wide. Gemmae lacking. In páramo bogs, very rare............................

..................... Heterogemma patagonica (Herzog & Grolle) L. Söderstr. & Váňa

5. Leaf cells smaller, 30-40 µm in midleaf. Stem epidermis cells less than 4× longer than wide. Gemmae presen.......... Schis-

tochilopsis incisa (Schrad.) Konstant.6. Trigones large. Leaf apex acute to

acuminate. Plants reddish-brown or purple, rarely green..... Anastrophyllum

240

6. Trigones lacking or small. Leaf apex obtuse to subacute. Plants green or brownish, rarely purplish (Sphenolobus austroamericanus)............................... 7

7. Leaf cells with conspicuous trigones, walls not uniformly thickened. Ventral stolons present. Gemmae absent.............

.......................................... Lophonardia7. Leaf cells with ± uniformly thickened

walls, trigones inconspicuous. Ventral stolons absent. Gemmae present or absent. Rare species of (super)páramo. 8

8. Leaves densily imbricate, leaf apices acute. Leaf cells 20-30 µm in midleaf. Plants brownish, with a strong resin smell, paroicous, usually copiously fertile. Gemmae reddish. In superpáramo above 4000 m, known from Ecuador, to be expected in Colombia........................

Isopaches bicrenatus (Hoffm.) H.Buch8. Leaves distant or laxly imbricate, leaf

apices acute or obtuse. Leaf cells smaller, 15-20 µm in midleaf. Plants dioicous, frequently sterile, without resin smell. Gemmae absent or reddish-brown....... 9

9. Leaves succubous, leaf apices obtuse. Plants greenish-brown, ca. 1 mm wide. Perianth eplicate, completely smooth.....

................................................... Gymnocoleopsis cylindriformis (Cephaloziellaceae)9. Leaves transverse, leaf apices (sub)acute.

Plants reddish or greenish-brown, 0.5-1 mm wide. Perianth plicate, sometimes with a white mouth........ sphenolobus

Key 11. Introductory key to Porellales (foliose liverworts II)

1. Underleaves present............................. 21. Underleaves lacking............................. 52. Lobule for most of its length attached to

the lobe along a keel. Keel and lobule usually widely spreading from stem. Plants variously colored but never red or purple............... Lejeuneaceae: Key 12

2. Lobule almost free from the lobe, keel lacking or indistict. Lobule parallel to the stem or weakly spreading............... 3

3. Underleaves undivided. Lobules plane, with rounded apex, margins of lobule often dentate to laciniate, at least near the base. Plants green to brown, robust, 3-7 mm wide.............................. Porella

3. Underleaves divided (rarely undivided). Lobules inflated, sac-like (or plane, then with acute apex), margins of lobule ± entire. Plants green to reddish or purple, 0.5-2.5 mm wide................................. 4

4. Plants pure green, without any trace of reddish pigmentation. Leaf margins toothed. Lobules very small, attached to the ventral margin of the lobe, at some distance from the stem...........................

...................... Jubula bogotensis Steph.4. Plants usually with reddish pigmentation,

rarely pure green. Leaf margins entire (rarely toothed). Lobules usually very close to the stem (at some distance from stem in the Diastaloba group: small reddish plants)........................ Frullania

5. Stems fragile, epidermis of only 5-6 rows of cells. Rhizoids from ventral surface of stem. Lobules very narrowly attached to stem (by only 1-4 cells). Plants very small, usually less than 1 mm wide. Leaf cells with many small, colorless oil bodies..................................

.......................... Lejeuneaceae: Key 125. Stems rigid, epidermis of numerous

thick-walled cells. Rhizoids in bundles from the surface of the lobule. Lobules more broadly attached to stem. Plants usually larger. Leaf cells with 1-2 large, brownish oil bodies.................... Radula

Key 12. Lejeuneaceae

1. Leaves highly specialized, distal part forming an inflated sac (= extension of the lobule)................................... Colura


241

Gradstein

1. Leaves less specialized, distal part not forming an inflated sac......................... 2

2. Underleaves lacking............................. 32. Underleaves present (sometimes very

small)................................................... 43. Plants growing upright from a

stoloniferous base, in or near running water. Stems rigid, of thick-walled cells. Known from Ecuador, to be expected in Colombia.................................... Myrio- coleopsis gymnocolea E.Reiner & Gradst.

3. Plants creeping, without stoloniferous base. Stems fragile, of thin-walled cells..

.......................................... Cololejeunea4. Underleaves undivided to weakly

emarginate............................... Key 12a 4. Underleaves bifid.................... Key 12b

Key 12a. Lejeuneaceae withundivided underleaves

1. Leaf margins toothed, at least near apex ............................................................. 21. Leaf margins entire............................ 152. Ventral merophyte 4 or more cells wide ............................................................. 3 2. Ventral merophyte 2(-3) cells wide.... 11 Width of the ventral merophyte is

measured by the number of epidermis cells across the ventral stem surface (= area between where the underleaves are attached, but not in the direct neighborhood of the underleaf base because there the number of epidermis cells is variable).

3. Leaf cells with evenly thickened walls, trigones lacking. Lobules with a very long, curved tooth (5-10 cells long). Underleaves toothed, emarginate.Fulfor- dianthus pterobryoides (Spruce) Gradst.

3. Leaf cells with trigones, walls not evenly thickened. Lobules with a shorter tooth (less than 5 cells long) or with several teeth (or without tooth). Underleaves entire or toothed................................... 4

4. Median leaf cells elongate; trigones cordate................................................. 5

4. Median leaf cells isodiametric, trigones various, not cordate............................. 8

5. Underleaves toothed............................ 65. Underleaves entire............................... 76. Plants pinnate or dichotomous,

branches predominantly Frullania-type. Innovations lacking.............. Bryopteris

6. Plants irregularly branched, branches predominantly Lejeunea-type. Innova-tions present................................ Thysa-

nanthus amazonicus (Spruce) Schiffn.7. Lobules with 7-9 teeth. Underleaf apex

± rounded, base auriculate. Perianth 8-10-keeled, with innovations. High Andes of Peru and Ecuador, to be expected in Colombia........... Frullanoi-

des laciniatiflora (Loitl.) van Slageren.7. Lobules with 1-3 teeth. Underleaf

apex emarginate, base not auriculate. Perianth 3-keeled, without innovations. Throughout tropical America.. Caudale-

jeunea lehmanniana (Gottsche) A. Evans8. Underleaf insertion line ± straight.

Plants 1.5-2 mm wide. Perianth with two ventral keels, innovations lacking.... Lo-

pholejeunea nigricans (Lindenb.) Schiffn.8. Underleaf insertion line deeply arched.

Plants usually more than 2 mm wide. Perianth without ventral keels, innovations present or absent............... 9

9. Apical portion of leaf broadly recurved. Ventral merophyte 4 cells wide, epidermal cells thin-walled. Plants glossy brown................................ Lindi-

gianthus cipaconeus Kruijt & Gradst.9. Leaf ± plane. Ventral merophyte more

than 4 cells wide, epidermal cells thick-walled................................................ 10

10. Lobules plane, with (1-)2-4 teeth. Perianth terminal on main stem or elongated branches, with two long innovations........................ Marchesinia

10. Lobules strongly inflated-rounded, small (hidden behind the underleaves),

242

± without teeth. Perianth on a very short branch, with one short innovation or without innovation..................................

........... Symbiezidium dentatum Herzog11. Underleaves very large, 6-10× stem

width, apex very short-bifid. Perianth terete, without keels................ Lejeunea

sulphurea (Lehm. & Lindenb.) Spruce11. Underleaves smaller, apex undivided

(emarginate in Harpalejeunea). Perianth with keels........................................... 12

12. Plants pale green. Disciform gemmae usually produced on dorsal leaf margins. Underleaf apex truncate... Cyclolejeunea

convexistipa (Lehm. & Lindenb.) A.Evans 12. Plants brownish green. Disciform

gemmae lacking................................. 13 13. Leaves with (5-)7-25 teeth. Underleaves

toothed or entire. On living leaves or bark ..................................... Odontolejeunea

13. Leaves with 1-5 teeth. Underleaves entire. On bark or rock....................... 14

14. Branches predominantly Frullania-type. Female bracteole toothed. Dicranolejeu-

nea axillaris (Nees & Mont.) Schiffn.14. Branches predominantly Lejeunea-type.

Female bracteole entire. Acanthocoleus aberrans (Lindenb. & Gottsche) Kruijt 15. Underleaves toothed............................ 615. Underleaves entire............................. 1616. Leaves with ocelli (scattered, in a row or

1-2 ocelli near leaf base).................... 1716. Leaves without ocelli......................... 2317. Ocelli in a row or 1-2 near leaf base.. 1817. Ocelli scattered.................................. 20 18. Row of ocelli 7-22 cells long. Ventral

merophyte 4-6 cells wide. Lobule tooth 3-5 cells long. Canopy epiphyte in low-land rainforest of Brazil; to be expec-ted in the Colombian Amazon. Neu-

rolejeunea seminervis (Spruce) Schiffn.18. Row of ocelli 1-6 cells long. Ventral

merophyte 2(-3) cells wide. Lobule tooth 1 cell long ................................ 19

19. Lobule fully inflated, bottle-shaped, with a long, falcate tooth. Perianth sharply

5-keeled, the keels smooth or toothed, never extending into inflated horns.........

....................................... Harpalejeunea19. Lobule inflated in the lower half,

flattened above, rectangular, with a short, blunt tooth. Perianth 4-keeled, the keels smooth and extended into inflated horns............................. Ceratolejeunea

20. Ventral merophyte 4 or more cells wide............................................................. 21

20. Ventral merophyte 2-3 cells wide (see note in couplet 2)............................... 22

21. Underleaf apex undivided. Frullania-type branches present. Leaf apex roun-ded ................................. Stictolejeunea

21. Underleaf apex emarginate. Frullania-type branches lacking. Leaf apex acute, rarely rounded............................... Lepi-

dolejeunea eluta (Nees) R.M.Schust.22. Leaf cells with large trigones. Ocelli

conspicuous, usually larger than green leaf cells. Epidermal cells thick-walled. Lobules curved downward............... Lu-

teolejeunea herzogii (Buchloh) Piippo22. Leaf cells with minute trigones. Ocelli

inconspicuous, equal in size to or smaller than other leaf cells. Epidermal cells thin-walled. Lobules straight or somewhat curved upwards........ Lepido-

lejeunea sullivantii (Gottsche) E.Reiner23. Ventral merophyte 4 or more cells wide

........................................................... 2423. Ventral merophyte 2 cells wide (see note

in couplet 2)....................................... 4724. Trigones ± cordate. Leaves convolute

when dry (rarely plane)...................... 2524. Trigones various but not cordate. Leaves

spreading when dry (rarely slightly convolute).......................................... 35

25. Underleaf apex emarginate................ 2625. Underleaf apex rounded or truncate, not

emarginate......................................... 2726. Plants dark green to brown to blackish,

rather robust, 2-3.5 mm wide, leaves when dry strongly convolute. Stems rigid, without hyalodermis, ventral


243

Gradstein

merophyte 8 cells wide. Perianth with innovations............................................. ................... Thysananthus amazonicus

26. Plants light green to pale yellowish-brown, smaller, 1.5-2 mm wide, leaves usually not convolute. Stems flaccid, with a distinct hyalodermis, ventral merophyte 4 cells wide. Perianth without innovations.............................................

................Caudalejeunea lehmanniana27. Lobules with 3-10 teeth (the teeth

sometimes inflexed)........................... 2827. Lobules with 0-2 teeth (occasionally 3

teeth: Mastigolejeunea innovans)...... 3028. Plants blackish in older stem portions.

Perianth with 5-10 keels, with inno-vations .............................. Frullanoides

28. Plants not blackish. Perianths with 3-10 keels, innovations present or absent.......

........................................................... 2929. Underleaf insertion line slightly curved,

underleaf base plane. Perianth with 5-10 keels, innovations lacking. Flagelliform branches (producing caducous leaves) frequently present. Lowlands (0-800 m)

......................................... Acrolejeunea29. Underleaf insertion line deeply arched,

underleaf bases folded. Perianth with 3(-4) keels, innovations present. Flagelliform branches lacking. Montane (300-2000 m).............. Brachiolejeunea

30. Ventral epidermis cells not or scarcely larger than medulla cells (stem cross-section). Medulla cells thick-walled. Plants brown-green to blackish-green. 31

30. Ventral epidermis cells distinctly larger than medulla cells. Medulla cells thin-walled................................................ 33

31. Leaf apex acute. Female bracts and bracteole toothed. Amazonia..................

................... Thysananthus amazonicus31. Leaf apex rounded. Female bracts and

bracteole without teeth....................... 3232. Plants turning olive- to reddish-brown.

Innovations lacking. Female bract apices acute-acuminate...................................... ............................... Schiffneriolejeunea

32. Plants turning dark-brown to black. Innovations present. Female bract apices rounded....................... Mastigolejeunea

33. Lobules mostly reduced.......................... ....................... Acanthocoleus aberrans33. Lobules well-developed, never reduced,

with two teeth.................................... 3434. First lobule tooth short, incurved and

blunt, second tooth long, pointing outwards, sharp. Lobule truncate. Leaves not squarrose when moist. Upper montane (above 2000 m)........................

.................................. Blepharolejeunea34. Lobule teeth equal or the first tooth

larger than the second. Lobule oblique or truncate. Leaves squarrose when moist. Lowland and montane (300-2000 m)........................................ Brachiolejeunea

35. Median leaf cells distinctly elongate, ca. 2 × longer than wide. [Plants robust. Ventral merophyte more than 10 cells wide. Lobules often reduced. Perianth with 5-8 keels]. In cacao plantations of western Ecuador, not yet known from Colombia...................................... Spru-

ceanthus theobromae (Spruce) Gradst.35. Median leaf cells isodiametrical or

slightly elongate................................. 3636. Lobules with 2-4 teeth....................... 3736. Lobules with 0-1 tooth....................... 4037. Leaf apex strongly and broadly recurved,

acute. Epidermal cells thin-walled. Plants glossy brown. Above 2000 m.......

........................................ Lindigianthus cipaconeus (Gottsche) Kruijt & Gradst.

37. Leaf apex ± plane, rounded or acute-acuminate. Epidermal cells thick-walled ........................................................... 38

38. Insertion line of underleaves deeply arched (more than 100 µm deep). Ventral merophyte 6-12 cells wide. Plants more than 2 mm wide, often black. Perianth without ventral keels......... Marchesinia

38. Insertion of underleaves straight or shallowly curved. Ventral merophyte 4 cells wide. Plants smaller, never black. Perianth with two ventral keels......... 39

244

39. Plants whitish, pale yellowish or grayish. Oil bodies 1-2(-3) per leaf cell, very large, filling up the cell lumen. Second lobule tooth much larger than the first tooth. Androecia on short-specialized branches, male bracteoles reduced..........

.. Cheilolejeunea (sect. Leucolejeunea)39. Plants pale green to brown. Oil bodies

more than 3 per leaf cell, rather small, not filling up the cell lumen. Lobule teeth identical or the first tooth larger than the second tooth. Androecia on elongate shoots, male bracteoles not reduced.............................. Dibrachiella

40. Underleaf insertion line deeply arched, over 100 µm deep............................... 41

40. Underleaf insertion line straight or shallowly curved................................ 42

41. Lobules less than 1/4 leaf length. Leaf cells with radiate trigones. Perianth on a very short branch (appearing lateral on the stem), keels ciliate-laciniate. Lowland and montane.... Symbiezidium

41. Lobules 1/4-1/2× leaf length. Leaf cells usually with large bulging trigones. Perianth on an elongate shoot, keels smooth. Exclusively montane.................

.. Cheilolejeunea (sect. Omphalanthus)42. Plants whitish, pale-yellowish or

grayish. Oil bodies 1-2(-3) per leaf cell, very large, filling up the cell lumen........

...Cheilolejeunea (sect. Leucolejeunea)42. Plants darker: green, brown or black. Oil

bodies more than 3 per leaf cell, smaller, not filling up the cell lumen............... 43

43. Midleaf cells small, 10-20 µm in dia-meter, trigones confluent, walls almost evenly thickened. Lobules flask-shaped, often darker than the leaf-lobe. Plants usually blackish............. Neurolejeunea

43. Midleaf cells larger, trigones not confluent, walls not evenly thickened. Lobules not flask-shaped, not darker than the leaf-lobe. Plants green, brown or black.............................................. 44

44. Epidermis cells distinctly larger than medulla cells (stem cross-section).

Perianth keels ciliate-laciniate, inno-vations lacking. Plants glossy black or dark-brown, rarely green. Oil bodies homogeneous................ Lopholejeunea

44. Epidermis cells not or little larger than medulla cells. Perianth keels smooth or denticulate, innovations present, rarely lacking. Plants green or brown. Oil bodies granular................................... 45

45. Innovations lacking. Free margin of lobule incurved, at least near apex. Leaves when moist obliquely spreading, ± squarrose. Amazonia Verdoornianthus

45. Innovations present. Free margin plane. Leaves when moist widely spreading, not squarrose. Widespread, common. 46

46. Lobules never reduced. Underleaves (3-)4-7× stem width, imbricate. Innovations pycnolejeuneoid (basal leaf on innovation is an underleaf: see Gradstein et al. 2001: Fig. 4I). Dioicous (rarely paroicous). Plants yellow-brown to dark brown.................. Archilejeunea

46. At least some lobules reduced. Underleaves 2-4× stem width, distant to subimbricate. Innovations lejeuneoid (basal leaf on innovation is a lateral leaf: see Gradstein et al. 2001: Fig. 4J). Autoicous. Plants pale-green to greenish-brown to black.... Dibrachiella

47. Lobules strongly swollen, ball-shaped, free margin inrolled 2-3 times. Anoplo-

lejeunea conferta (Meissn.) A. Evans47. Lobules not ball-shaped, free margin

plane or slightly incurved.................. 4848. Frullania-type branches present

(sometimes only few). Lobules when well-developed with 2 teeth. Plants greenish-brown.................................. 14

48. Frullania-type branches lacking. Lobules with 1 tooth. Plants pale green... ........................................................... 49

49. Plants tiny, 0.4-0.7 mm wide. Leaf lobes suberect to obliquely spreading. Lobule tooth long-acute, consisting of a strongly elongate cell (at least 4× as long as wide). Hyaline papilla distal to the


245

Gradstein

tooth. Underleaves very small, 1.5-2× stem width, orbicular..............................

........................................ Cheilolejeuna holostipa (Spruce) Grolle & R.L.Zhu

49. Plants larger. Leaves wide-spreading. Lobule tooth blunt or acute, short. Hyaline papilla proximal to the tooth (except in L. herminieri (Steph.) R.L.Zhu). Underleaves larger.. Lejeunea

Key 12b. Lejeuneaceae with bifid underleaves

1. Leaves highly specialized, upper part forming an inflated sac............... Colura

1. Leaves less specialized, upper part not forming an inflated sac......................... 2

2. Plants densely pinnate, growing in or near running water, with numerous short sexual branches. Stems robust, ventral merophyte (4-)6-14 cells wide. Known from Ecuador and Brazil, to be expected in Colombia......................................... 3

2. Plants not densely pinnate. Stems thinner, ventral merophyte 2(-6) cells wide ..................................................... 4

Width of the ventral merophyte is measured by the number of epidermis cells across the ventral stem surface (= area between where the underleaves are attached, but not in the direct neighborhood of the underleaf base because there the number of epidermis cells is variable).

3. Plants growing upright or pendent from a rhizome-like base. Lobules lacking on upright or pendent stems. Leaves transversely inserted, insertion line very short. Underleaves minute. Paroicous; antheridia in axils of female bracts.........

Colura irrorata (= Myriocolea irrorata)3. Plants creeping or pendent, without

rhizome-like base. Lobules present. Leaves longitudinally inserted, insertion line long. Underleaves well-developed. Autoicous; antheridia on separate

branches.................................................. .. Lejeunea (subg. Neopotamolejeunea)

4. Underleaf lobes conspicuously diverging (spreading outwards)........................... 5

4. Underleaf lobes not conspicuously diverging............................................ 13

5. One underleaf to each leaf, the underleaves usually densely overlapping. Leaf apex rounded...... Diplasiolejeunea

5. One underleaf to every alternate leaf, the underleaves usually distant. Leaf apex acute-acuminate to rounded................. 6

6 Underleaf lobes mostly 1 cell wide (except at the base), apex acute............ 7

6. Underleaf lobes broader, apex blunt... 107. Leaf apex rounded............................... 87. Leaf apex acute or acuminate.............. 98. Lobule tooth short and blunt, not falcate.

Ocellus near leaf base very large, 2.5-4× longer than adjacent leaf cells. Gynoecium without innovation.............. ........................................ Leptolejeunea

8. Lobule tooth elongate, falcate. Ocellus near leaf base smaller, maximally 2× longer than adjacent leaf cells. Gynoecium with innovation. In lowland rainforest................... Drepanolejeu-nea polyrhiza (Nees) Grolle & R.L.Zhu

9. Ocellus near leaf base very large, 2.5-4× longer than surrounding leaf cells. Lobule tooth short and blunt, not falcate. Gynoecium without innovation.............. ........................................ Leptolejeunea

9. Ocellus at leaf base smaller, 1-2× longer than surrounding leaf cells, or lacking. Lobules with a long, falcate tooth (rarely short and blunt). Gynoecium usually with innovation (rarely without)............. ................................... Drepanolejeunea

10. Leaf lobes with an unbroken row of 4-8 ocelli, ocelli usually glossy yellowish-brown. In the outer canopy of Amazonian rainforest....... Cheilolejeunea urubuensis (Zartman & I.L.Ackerman) Wei et al.

10. Leaf lobes with a broken or unbroken row of ( 1-)2-3 ocelli in the lower half,

246

the ocelli colorless or grayish. Lowland and montane....................................... 11

11. Ocelli in a broken row............................ ................................... Drepanolejeunea

11. Ocelli in an unbroken row.................. 1212. Innovations pycnolejeuneoid (basal

leaf on innovation is an underleaf; see Gradstein et al. 2001: Fig. 4I). Cells on dorsal leaf surface mamillose, thick-walled, with a broad, lens-shaped papilla. Lobule with or without pre-apical tooth (= tooth situated at the distal end of the lobule, at the junction with the ventral leaf margin)............... Drepanolejeunea

12. Innovations lejeuneoid (basal leaf on innovation is a lateral leaf; see Gradstein et al. 2001: Fig. 4G). Cells on dorsal leaf surface smooth, thin-walled (rarely thick-walled and with a conical papilla: H. scabra). Lobule without pre-apical tooth .............................. Harpalejeunea

13. Underleaves very large, at least 6× wider than the stem...................................... 14

13. Underleaves smaller........................... 1514. Leaf cells uniformly thin-walled or with

small trigones. Lobules often reduced; hyaline papilla proximal to the tooth at the lobule apex (see Gradstein et al. 2001: Fig. 4G). Plants glistening pale green, yellowish-green or whitish (stems often long and slender, pendent). Oil bodies small, numerous per cell, finely granular or homogeneous....... Lejeunea

14. Leaf cells with conspicuous trigones. Lobules not reduced; hyaline papilla distal to the tooth at the lobule apex (see Gradstein et al. 2001: Fig. 4H). Plants dull pale green to olive green. Oil bodies very large, 1-3 per cell, sausage-shaped, coarsely granular.....................................

...................................... Cheilolejeunea15. Lobules when well-developed very

large, 3/5-4/5 of lobe length. Leaves usually suberect, almost parallel to the stem. Plants minute, 0.2-0.6 mm wide............................................................... 16

15. Lobules smaller. Leaves spreading. Plants small or large........................... 17

16. Plants monoicous. Leaves without ocelli. Innovations pycnolejeuneoid (basal leaf on innovation is an underleaf; see Gradstein et al. 2001: Fig. 4I)..........

.................................................. Metale- jeunea cucullata (Reinw. et al.) Grolle16. Plants dioicous. Leaves usually with 1 or

more ocelli (at the base or scattered in the leaf). Innovations lejeuneoid (basal leaf on innovation is a lateral leaf; see Gradstein et al. 2001: Fig. 4J).................

........................................ Microlejeunea17. Ocelli present in leaves...................... 1817. Ocelli lacking in leaves...................... 3018. Ocelli present in underleaves............. 1918. Ocelli lacking in underleaves............. 2119. Ocelli reddish or brown. Leaf cells with

a broad papilla or smooth. Caducous leaves absent. Gynoecia on a very short branch, without innovations....................

......................................... Pictolejeunea19. Ocelli colorless or gray. Leaf cells

smooth. Caducous leaves present or absent. Gynoecia on long shoots, with 1-2 innovations.................................. 20

20. Caducous leaves present, produced on flagelliform branches............... Rectole- jeunea emarginiflora (Schiffn.) A.Evans

20. Caducous leaves absent or present, when present not produced on flagelliform branches........................ Lepidolejeunea

21. Underleaf lobes broadly rounded. [Ocelli 2-3 in an unbroken row in the lower half of the leaf]........................ 12

21. Underleaf lobes (sub)acute to acuminate........................................... 22

22. Leaves with a very large ocellus near the base, the basal ocellus 2.5-4× larger than adjacent leaf cells. Underleaf lobes subulate (1-2 cells wide), inserted at the outer edges of the underleaf. Underleaf margins bordered by large cells............................. Leptolejeunea


247

Gradstein

22. Basal ocellus smaller, maximally 2× larger than adjacent leaf cells. Underleaf lobes wider, not inserted at outer edges of underleaf. Underleaf margins not bordered by large cells....................... 23

23. Plants brown or brownish-green. Walls of leaf cells ± pale brown, trigones radiate (or lacking). Branch bases sometimes with a huge, strongly inflated lobule (“utricle”). Perianth usually with inflated horns (horns sometimes absent...

...................................... Ceratolejeunea23. Plants green or whitish (rarely brown).

Walls of leaf cells colorless, trigones not radiate. Utricles lacking. Perianth without inflated horns........................ 24

24. Leaf lobes with a long, unbroken row of 4-8 ocelli, the ocelli usually glossy yellowish-brown. In the canopy of Amazonian rainforest.............................

................... Cheilolejeunea urubuensis24. Leaf lobes not with an unbroken row of

4-8 ocelli. Lowland and montane...... 2525. Underleaf margins denticulate to

crenulate to subentire. Gemmae usually present on leaf margins. Perianth flattened, 2-keeled, the keels expanded above into short auricles. On living leaves, occasionally on bark or rock.......

........................................ Cyclolejeunea25. Underleaf margins entire (sometimes

with a small tooth: Xylolejeunea). Gemmae lacking. Perianth 3-5-keeled in the upper half, the keels not expanded into auricles. On bark or rotten wood.. ............................................................. 26

26. Caducous leaves present, very small, much smaller than ordinary leaves, produced on flagelliform branches. Leaf cells very small, 14-18 µm long in the middle of the lobe, walls thin or uniformly thickened, without trigones. Lobules usually with a long, falcate tooth. [Plants less than 1 mm wide; leaves wide-spreading]......................................

........................................ Rectolejeunea

26. Caducous leaves present or absent, when present not smaller than ordinary leaves. Leaf cells larger. Lobule tooth short, not falcate ................................................ 27

27. Plants very small, less than 0.7 mm wide. Leaves suberect or obliquely spreading. Ocelli grayish or brown..Microlejeunea

27. Plants larger. Leaves wide-spreading. Ocelli colorless or grayish................. 28

28. Epidermis cells thin-walled. Lobules sometimes reduced. Underleaves distant. Perianth with flat, antler-like lacinia near apex........................................................

........ Otigoniolejeunea huctumalcensis (Lin denb. & Gottsche) Y.M.Wei et al.28. Epidermis cells thick-walled. Lobules

never reduced. Underleaves distant or imbricate. Perianth without antler-like lacinia................................................ 29

29. Underleaves distant. Leaf cells thin-walled, without or with very small trigones. Leaf base with two transversal rows of larger, elongate cells, the upper, suprabasal row consisting of 2-5 adjacent ocelli................... Xylolejeunea

29. Underleaves (sub)imbricate. Leaf cells with conspicuous trigones. Leaf base not with two transversal rows of elongate cells, the ocelli 2-20 per leaf, adjacent or scattered......................... Pycnolejeunea

30. Leaf margins (and sometimes the entire dorsal leaf surface) toothed by conically projecting cells with thick-walled tips ... ........................................................... 31

30. Leaf margins entire or ± toothed, but not toothed by conically projecting cells with thick-walled tips........................ 32

31. Dorsal leaf surface spinose. Perianth inflated, with 5 keels................ Lejeunea

31. Dorsal leaf surface smooth or papillose, not spinose. Perianth flattened, with 2 keels .............................. Prionolejeuna

32. Vegetative reproduction by means of large, multicellular, rounded or elongate gemmae from leaf margins. [Plants usually growing on living leaves.

248

Margins of leaves and underleaves ± toothed] .......................... Cyclolejeunea

32. No vegetative reproduction by means of multicellular gemmae........................ 33

33. Vegetative reproduction by means of caducous leaves................................. 34

33. No vegetative reproduction by caducous leaves................................................. 36

34. Leaf cells small, less than 20 µm in diameter, trigones lacking. Caducous leaves much smaller than ordinary leaves, usually produced on special fla-gelliform branches.......... Rectolejeunea

34. Leaf cells larger, trigones present or absent. Caducous leaves not smaller than ordinary leaves, special flagelliform branches usually lacking.................... 35

35. Lobule tooth long and sharp, hyaline papilla distal to the tooth. Lobules not reduced...................................................

..Cheilolejeunea adnata (Kunze) Grolle35. Lobule tooth short and blunt, hyaline

papilla proximal to the tooth. Lobules sometimes reduced................. Lejeunea

36. All or part of the leaves conspicuously elongate, (1.2-)2-4 times longer than wide ................................................... 37

36. Leaves less elongate.......................... 4037. Leaves suberect, apex acute-acuminate.. ................................... Drepanolejeunea37. Leaves obliquely to widely spreading,

apex rounded...................................... 3838. Leaves with very short insertion,

attached to the stem by only 2-3 cells. Lobules ± ligulate, flat, standing upright almost parallel to the stem, keel very short. Plants rheophytic (growing in rivers). Amazonia, not yet known from Colombia.................................... Schus- terolejeunea inundata (Spruce) Grolle

38. Leaves with a longer insertion. Plants not rheophytic.................................... 39

39. In wet leaf axils of tank bromeliads. Midleaf cells in elongate leaves 35-70 µm long, 1.5-4× longer than wide. Perianth inflated, 5-keeled, keels not expanded into auricles............................

............. Bromeliophila helenae Gradst.39. On living leaves in lowland rainforest.

Midleaf cells smaller, 20-30 µm long, subisodiametrical. Perianth flattened, 2-keeled, keels expanded into large auricles....... Otolejeunea schellii Tixier

40. Walls of leaf cells ± pale brown, trigones radiate (or lacking). Branch bases sometimes with a huge, strongly inflated lobule (= utricle). Perianth usually with horns. Plants brown or brownish-green..

...................................... Ceratolejeunea40. Walls of leaf cells colorless, trigones

not radiate. Utricles lacking. Perianth without horns. Plants green or whitish, rarely greenish-brown........................ 41

41. Trigones large or small. Lobules never reduced, always strongly inflated. Oil bodies large (more than 1/2 the cell lumen in length), coarsely segmented. Hyaline papilla distal to the lobule tooth (or between 2 small, closely associated teeth at the leaf apex) (see Gradstein et al. 2001: Fig. 4H).......... Cheilolejeunea

41. Trigones lacking or very small. Lobules sometimes reduced, inflated or not inflated. Oil bodies smaller (less than 1/4 the cell lumen in length), finely segmented or smooth. Hyaline papilla proximal to the lobule tooth (see Gradstein et al. 2001: Fig. 4G)...............

................................................ Lejeunea

LITERATURE CITED

BISCHLER, H., S.R. GRADSTEIN, S. JOvET-AST, D.G. LONG & N. SALAZAR ALLEN. 2005. Marchantiidae. Flora Neotropica Monograph 97: 1-262.

CAMPOS-SALAZAR, L. & J. URIBE-M. 2012. Taxonomic revision of Balantiopsaceae (Marchantiophyta) of Colombia. Hedwigia 94: 94-127.

COSTA, D.P. 2008. Metzgeriaceae (Hepaticae). Flora Neotropica Monograph 102: 1-170.

CRANDALL-STOTLER, B.J., R.E. STOTLER & D.G. LONG. 2009. Phylogeny and classification of the Marchantiophyta. Edin burgh Journal of Botany 66: 155-198.


249

Gradstein

DAUPHIN, G. 2003. Ceratolejeunea. Flora Neotropica Monograph 90: 1-86.

FREy, W. (ed.). 2009. Syllabus of plant families. Part 3. Bryophytes and seedless vascular plants. Borntraeger, Berlin, Stuttgart. 419 pp.

GIL-NOvOA J.E., M.E. MORALES-PUENTES & S.R. GRADSTEIN. 2015. Platycaulis reni folia R.M.Schust. In: L.T. Ellis (ed.). New national and regional bryophyte records. Journal of Bryology 37: 319.

GRADSTEIN, S.R. 2016. The genus Plagiochila (Marchantiophyta) in Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 40: 104-136.

GRADSTEIN, S.R. & A.L. ILkIU-BORGES. 2015. A taxonomic monograph of the genus Odontoschisma (Marchantiophyta: Cepha lo-zia ceae). Nova Hedwigia 100: 15-100.

GRADSTEIN, S.R. & J. URIBE-M. 2016. Marchantiophyta. In: R. Bernal, S.R. Gradstein & M. Celis (eds.), Catálogo de plantas y líquenes de Colombia: 282-352. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá.

GRADSTEIN, S.R., S.P. CHURCHILL & N. SALAZAR ALLEN. 2001. Guide to the Bryophytes of Tropical America. Memoirs of the New York Botanical Garden 86: 1-577.

GRADSTEIN, S.R., T. PóCS & R.L. ZHU. 2016. Lejeuneaceae. In: L. Söderström, A. Hagborg & M. von Konrat (eds). World checklist of hornworts and liverworts: 297-414. Phytokeys 59.

ILkIU-BORGES, A.L. 2016. Prionolejeunea (Lejeu-neaceae, Jungermanniopsida). Flora Neo tro pica Monograph 116: 1-131.

JUáREZ-MARTíNEZ, C., H. OCHOTERENA & C. DEL-GADILLO-MOyA. 2016. Cladistic analysis of the Stephaniellaceae (Marchantiophyta) based on morphological data. Systematics and Biodi-versity 14: 74-87.

MCNEILL, J., F.R. BARRIE, W.R. BUCk, v. DEMOULIN, W. GREUTER, D.L. HAWkSWORTH, P.S. HERENDEEN, S. kNAPP, k. MARHOLD, J. PRADO, W.F. PRUD’HOMME vAN REINE, G.F. SMITH, J.H. WIERSEMA & N.J. TURLAND. 2012. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Vegetabile 154: 1-208.

REINER-DREHWALD, M.E. & A. GODA. 2000. Revision of the genus Crossotolejeunea (Lejeuneaceae, Hepaticae). Journal of the Hattori Botanical Laboratory 89: 1-54.

SCHUSTER, R.M. 2002. Austral Hepaticae II. Nova Hedwigia Beiheft 119: 1-606.

SHI X-Q., S.R. GRADSTEIN & R.L. ZHU. 2015. Phylogeny and taxonomy of Archilejeunea (Spruce) Steph. (Lejeuneaceae subfam. Ptychanthoideae, Marchantiophyta) based on molecular markers and morphology. Taxon 64: 881-892.

SöDERSTRöM L., A. HAGBORG, M. vON kONRAT, S. BARTHOLOMEW-BEGAN, D. BELL, L. BRISCOE, E. BROWN, D.C. CARGILL, D.P. COSTA, B.J. CRANDALL-STOTLER, E.D. COOPER, G. DAUPHIN, J.J. ENGEL, k. FELDBERG, D. GLENNy, S.R. GRADSTEIN, X.L. HE, J. HEINRICHS, J. HENTSCHEL, A.L. ILkIU-BORGES, T. kATAGIRI, N.A. kONSTANTINOvA, J. LARRAIN, D.G. LONG, M. NEBEL, T. PóCS, F. FELISA PUCHE, E. REINER-DREHWALD, M.A.M. RENNER, A. SASS-GyARMATI, A. SCHAFER-vERWIMP, J.G.S. MORAGUES, R.E. STOTLER, P. SUkkHARAk, B.M. Thiers, J. Uribe-M., J. Váňa, J.C. Villarreal, M. WIGGINTON, L. ZHANG & R.L. ZHU. 2016. World checklist of hornworts and liverworts. PhytoKeys 59: 1-821.

STOTLER, R., B.C. CRANDALL-STOTLER & C. CARGILL. 2016. Fossombroniaceae. In: L. Söderström, A. Hagborg & M. von Konrat (eds.). World checklist of hornworts and liverworts: 463-467. Phytokeys 59.

URIBE-M., J. 2008. Monografía del subgénero Meteoriopsis del género Frullania (Frullaniaceae, Marchantiopsida). Caldasia 30: 49-94.

URIBE-M., J. & J. AGUIRRE-C. 1997. Clave para los géneros de hepáticas de Colombia. Caldasia 19: 13-27.

Váňa, J. 2003. Notes on Gymnomitriaceae (subf. Gymnomitrioidae) in Latin America. Acta Academiae Paedagogiae Agriensis, Sectio Biologiae 24: 109-128.

yE, W., S.R. GRADSTEIN, J.A. SHAW, B. SHAW, B.C. HO, A. SCHäFER-vERWIMP, T. PóCS, J. HEINRICHS & R.L. ZHU. 2015. Phylogeny and classification of Lejeuneaceae subtribe Cheilolejeuneinae (Marchantiophyta) based on nuclear and plastid molecular markers. Cryptogamie, Bryologie 36: 313-333.

yU, y., T. PóCS & R.L. ZHU. 2015. Notes on Early Land Plants Today. 62. A synopsis of Myriocoleopsis (Lejeuneaceae, Marchantio-phyta) with special reference to transfer of Cololejeunea minutissima to Myriocoleopsis. Phytotaxa 183: 293-297.

Recibido: 17/08/2016Aceptado: 25/10/2016

a new key to the genera of liverworts of colombia · dumort., clevea lindb., isopaches buch,...

Documents